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Some well-managed organic soils are known to have higher crop yield potential than conventionally managed soils due to the greater soil quality and the ability to tolerate weed competition. However, low available soil mineral N and P in some organic systems may mask such soil quality–related benefits. We hypothesize that when plant-available N and P are not limiting, tillage-based highly diverse organic crop rotations have less yield loss (better crop tolerance) due to weed competition and higher crop yields than no-till conventional systems with low-diversity rotations. A greenhouse study was carried out in Saskatoon, Canada, using long-term (18-yr) organically managed soils (ORG) and no-till conventional soils (CONV) with three crop rotation diversities (LOW, MEDIUM, and HIGH) to compare the crop tolerance to weed competition under standard soil nutrient management conditions and under excess supply of mineral N and P. Under fertilized conditions, crop biomass increased by 50% and 69% in ORG and CONV systems, respectively. Weed biomass was similar between ORG and CONV systems under nonfertilized conditions but was 14% greater in CONV when excessive N and P were supplied. Crop biomass loss (crop tolerance) was not different among cropping systems under excess fertilizer or under standard fertilizer levels. Even with greater weed biomass under fertilized conditions, the CONV system showed crop tolerance similar to that of the ORG system. Under nonfertilized conditions, the crop biomass yield was 43% lower in ORG compared with CONV, and even after mineral N and P were applied, ORG systems showed less (17%) crop biomass than CONV. Further, differences in crop tolerance were not identified among crop rotations under both fertilizer levels. Overall, this study revealed that there were no yield benefits or better crop tolerance to weed competition in organically managed soils compared with no-till conventional soils, even under nonlimiting soil macronutrient conditions.

Phenotyping crop performance is critical for line selection and variety development in plant breeding. Canola (Brassica napus L.) flowers, the bright yellow flowers, indeterminately increase over a protracted period. Flower production of canola plays an important role in yield determination. Yellowness of canola petals may be a critical reflectance signal and a good predictor of pod number and, therefore, seed yield. However, quantifying flowering based on traditional visual scales is subjective, time-consuming, and labor-consuming. Recent developments in phenotyping technologies using Unmanned Aerial Vehicles (UAVs) make it possible to effectively capture crop information and to predict crop yield via imagery. Our objectives were to investigate the application of vegetation indices in estimating canola flower numbers and to develop a descriptive model of canola seed yield. Fifty-six diverse Brassica genotypes, including 53 B. napus lines, two Brassica carinata lines, and a Brassica juncea variety, were grown near Saskatoon, SK, Canada from 2016 to 2018 and near Melfort and Scott, SK, Canada in 2017. Aerial imagery with geometric and radiometric corrections was collected through the flowering stage using a UAV mounted with a multispectral camera. We found that the normalized difference yellowness index (NDYI) was a useful vegetation index for representing canola yellowness, which is related to canola flowering intensity during the full flowering stage. However, the flowering pixel number estimated by the thresholding method improved the ability of NDYI to detect yellow flowers with coefficient of determination (R2) ranging from 0.54 to 0.95. Moreover, compared with using a single image date, the NDYI-based flowering pixel numbers integrated over time covers more growth information and can be a good predictor of pod number and thus, canola yield with R2 up to 0.42. These results indicate that NDYI-based flowering pixel numbers can perform well in estimating flowering intensity. Integrated flowering intensity extracted from imagery over time can be a potential phenotype associated with canola seed yield.

Silene latifolia Poiret of Eurasia has established in North America, prompting this structural study of its mature unisexual buds and flowers. Floral nectaries, anther and stigma changes, and vestigial reproductive structures were studied using light and scanning electron microscopy. In staminate flowers, anthers dehisced before anthesis and >90% of their pollen was liberated within 36 h. Accumulated in the tubular calyx, nectar descended an anthophore from the stomatal-bearing nectary at the stamen bases. Nectary tissue surrounded the pistillode, a central filamentous organ lacking ovules but tipped by hairs resembling stigmatic papillae. In pistillate flowers, nectar flowed into an inflated calyx. The annular nectary had 10 regularly spaced, stomatal-lined craters and was continuous with the adaxial surfaces of the infertile antisepalous and epipetalous staminodes. Key elements of entomophilous pollination leading to successful sexual reproduction of this invasive species include secondary nectar presentation from disparate floral nectaries that, for pistillate flowers, also incorporate the staminodes; rapid pollen release from anthers; and elongation of papillae by tip growth that enhances each stigma’s receptive surface. Context is also provided for future studies of floral nectary development in this model dioecious species.

With the increasing resistance of wild oat (Avena fatua L.) to herbicides, there is a need to evaluate the potential of alternative cropping systems based on integrated weed management principles. A 5-yr field study at eight sites across Canada was used to evaluate the profitability of alternative cropping systems that have the potential to control wild oat using cultural practices in conjunction with herbicides. Cultural practices included twice the recommended seeding rates, fall-seeded winter crops, barley (Hordeum vulgare L.) silage, fallow, and alfalfa (Medicago sativa L.). Seven of the 14 cropping systems in this study did not include wild oat herbicide for three consecutive years, controlling wild oat entirely by cultural practices. Cropping system profitability varied by location. For many locations, combinations of barley silage and fall-seeded winter crops without wild oat herbicide application were as profitable as a system of canola (Brassica napus L.) and wheat (Triticum aestivum L.) with wild oat herbicide applied every year. Unprofitable systems generally included those with fallow, alfalfa, and fall-seeded winter crops in regions with rates of high winter kill. Wild oat control can be achieved with diverse cropping systems that are as profitable as conventional annual cropping that relies on herbicide control of wild oat.

Traditional methods of estimating frost damage to crops are labor-intensive and time-consuming. Remote sensing imagery and vegetation indices can be used for condition assessment, however, the util...

Interrow cultivation is a selective, in-crop mechanical weed control tool that has the potential to control weeds later in the growing season with less crop damage compared with other incrop mechanical weed control tools. To our knowledge, no previous research has been conducted on the tolerance of narrow-row crops to interrow cultivation. The objective of this experiment was to determine the tolerance of field pea and lentil to interrow cultivation. Replicated field experiments were conducted in Saskatchewan, Canada, in 2014 and 2015. Weekly cultivation treatments began at the 4-node stage of each crop, continuing for 6 wk. Field pea and lentil yield linearly declined with later crop stages of cultivation. Cultivating multiple times throughout the growing season reduced yield by 15% to 30% in both crops. Minimal yield loss occurred when interrow cultivation was conducted once at early growth stages of field pea and lentil; however, yield loss increased with delayed and more frequent cultivation events.Nomenclature: Field pea; Pisum sativum L.; lentil; Lens culinaris L.

The escalating evolution of weed species resistant to acetolactase synthase (ALS)-inhibitor herbicides makes alternative weed control strategies necessary for field crops that are dependent on this herbicide group. A fully integrated strategy that combined increased crop seeding rates (2X or 4X recommended), mechanical weed control with a minimum-tillage rotary hoe, and reduced-rate non—ALS inhibitor herbicides was compared with herbicides, rotary hoe, and seeding rates alone as a method of controlling ALS inhibitor—tolerant Indian mustard as a model weed. The full-rate herbicide treatment had the lowest weed biomass (98% reduction) and the highest yield of all treatments in 3 of 4 site-years, regardless of seeding rate. The fully integrated treatment at the 4X seeding rate had weed suppression rates equal to the full herbicide treatment at the recommended seeding rate. The fully integrated and reduced-rate herbicide treatments at the 4X seeding rate reduced weed biomass by 89% and 83%, respectively, compared with the control at the recommended seeding rate. The rotary hoe treatment alone resulted in poor weed control (≤38%), even at the highest seeding rate. Fully integrated and reduced-rate herbicide treatments at 2X and 4X seeding rates had yields equal to those of the full herbicide treatment at the recommended seeding rate. Partially or fully integrated weed control strategies that combine increased crop seeding rates and reduced-rate non—ALS inhibitor herbicides, with or without the use of a rotary hoe, can control weeds resistant to ALS-inhibitor herbicides, while maintaining crop yields similar to those achieved with full-rate herbicides. However, combining increased seeding rate, reduced-rate herbicides, and mechanical rotary hoe treatment into a fully integrated strategy maximized weed control, while reducing reliance on and selection pressure against any single weed control tactic.Nomenclature: Indian mustard, Brassica juncea (L.) Czern.

Concern over the development of herbicide-resistant weeds has led to interest in integrated weed management systems that reduce selection pressure by utilizing mechanical and cultural weed control practices in addition to herbicides. Increasing crop seeding rate increases crop competitive ability and thus can enhance herbicide efficacy. However, it is unknown how increasing the seeding rate affects an herbicide’s efficacy. The objective of this study was to examine the interaction between increasing seeding rate and herbicide dose to control weeds. To meet this objective, the herbicide fluthiacet-methyl was applied to field-grown lentil, with Indian mustard, a proxy for wild mustard, used as a model weed. The experiment was a factorial design with four lentil seeding rates and seven herbicide rates. Overall the herbicide dose response was altered by changing lentil seeding rate. Increasing lentil seeding rate decreased the weed biomass production when herbicides were not applied. In two of the four site-years, increasing lentil seeding rate lowered the herbicide ED50, the dose required to result in a 50% reduction in weed biomass. Increasing the crop seeding rate altered the dose response to provide greater weed control at lower herbicide rates compared with normal crop seeding rates. Increased seeding rates also resulted in higher and more stable crop seed yields across a wider range of herbicide dosages. These results suggest that dose–response models can be used to evaluate the efficacy of other weed management practices that can interact with herbicide performance.

Canola (Brassica napus L.) often experiences high seed losses prior to and during harvest operations. Pod drop and pod shatter are responsible for pre-harvest seed losses in canola although relatively little is known about the significance of pod drop to total pre-harvest seed losses in canola. This research describes a method to measure the predisposition for pod drop among canola genotypes that was developed and tested across different sites from 2013 to 2015. This method, referred to as pod-retention resistance, consists of the use of a force gauge to quantify the force required to break the pod pedicle at the pedicle-rachis junction. Two sets of field experiments were used, the first developed and modeled the factors that influence pod-retention resistance, pod drop and seed shatter. A second set of experiments was used to validate these findings on a broader range of genotypes and determine whether a relationship between pod retention resistance and pod drop could be determined. Results showed that the pod-retention resistance was significantly greater for pods on the proximal half of the rachis compared to the distal half of the rachis. Several factors contributed strongly to variation in pod retention resistance (rachis position, genotype and site) and pod drop (site, rachis position and genotype). The relative importance of individual factors varied between pod retention resistance and pod drop suggesting no relationship between the two. However, weight-adjusted pod retention resistance, i.e., average pod retention resistance corrected for the average weight of dropped pods, was strongly correlated with pod drop, particularly in the validation experiments. This method could be used by canola breeders to select for and develop genotypes with reduced predisposition for seed losses via pod drop.

Alternative cropping systems are gaining attention throughout the world in order to increase the sustainability of agro-ecosystems. On the Canadian prairies, the tillage-based crop-fallow system has been replaced by no-till reduced input systems or tillage-based organic systems with more diversity in crop rotations but with no external inputs. However, the long-term effects of these alternative systems on weed and yield parameters have not been assessed. A study to examine weed and crop yield dynamics under diverse cropping systems was conducted within a 18-year cropping systems study near Scott, Saskatchewan. The trial was laid out in a split-split-plot design with four replicates. The main plots were three levels of inputs; a high input system (HIGH) that used tillage and inputs to maximize yield, a reduced system (RED) that used no-till practices and minimal inputs, and a tillage-based organic system (ORG) with no external inputs. The subplots were cropping diversity (rotations); fallow-annual grains (LOW), diversified annual grains (DAG), and diversified annuals and perennial forage (DAP). The sub-sub plots were the six phases of each rotation. There was an input by rotation interaction for weed biomass but not for weed density and crop yields. ORG systems had 7× and 4× greater weed density (107 plants m −2 ), 4× higher weed biomass (154 kg ha −1 ), and 32% and 35% lower yields (1052 kg ha −1 ) than RED and HIGH systems respectively. RED and HIGH input systems had similar crop yields and lower weed density than ORG. The LOW rotation had the lowest weed density. LOW and DAG rotations had similar yields, which were higher than in DAP. All systems showed an increase in weed density and biomass over time but did not impact on crop yields which was increasing over time likely influenced by a concurrent increase in rainfall. This study concludes that eliminating tillage and reducing agrochemicals is possible but eliminating agrochemicals requires better crop rotations for weed management as well as for nutrient management.

In western Canada, more money is spent on wild oat herbicides than on any other weed species, and wild oat resistance to herbicides is the most widespread resistance issue. A direct-seeded field experiment was conducted from 2010 to 2014 at eight Canadian sites to determine crop life cycle, crop species, crop seeding rate, crop usage, and herbicide rate combination effects on wild oat management and canola yield. Combining 2× seeding rates of early-cut barley silage with 2× seeding rates of winter cereals and excluding wild oat herbicides for 3 of 5 yr (2011 to 2013) often led to similar wild oat density, aboveground wild oat biomass, wild oat seed density in the soil, and canola yield as a repeated canola–wheat rotation under a full wild oat herbicide rate regime. Wild oat was similarly well managed after 3 yr of perennial alfalfa without wild oat herbicides. Forgoing wild oat herbicides in only 2 of 5 yr from exclusively summer annual crop rotations resulted in higher wild oat density, biomass, and seed banks. Management systems that effectively combine diverse and optimal cultural practices against weeds, and limit herbicide use, reduce selection pressure for weed resistance to herbicides and prolong the utility of threatened herbicide tools.

The traditional visual rating system is labor-intensive, time-consuming, and prone to human error. Unmanned aerial vehicle (UAV) imagery-based vegetation indices (VI) have potential applications in high-throughput plant phenotyping. The study objective is to determine if UAV imagery provides accurate and consistent estimations of crop injury from herbicide application and its potential as an alternative to visual ratings. The study was conducted at the Kernen Crop Research Farm, University of Saskatchewan in 2016 and 2017. Fababean ( Vicia faba L.) crop tolerance to nine herbicide tank mixtures was evaluated with 2 rates distributed in a randomized complete block design (RCBD) with 4 blocks. The trial was imaged using a multispectral camera with a ground sample distance (GSD) of 1.2 cm, one week after the treatment application. Visual ratings of growth reduction and physiological chlorosis were recorded simultaneously with imaging. The optimized soil-adjusted vegetation index (OSAVI) was calculated from the thresholded orthomosaics. The UAV-based vegetation index (OSAVI) produced more precise results compared to visual ratings for both years. The coefficient of variation (CV) of OSAVI was ~1% when compared to 18-43% for the visual ratings. Furthermore, Tukey’s honestly significance difference (HSD) test yielded a more precise mean separation for the UAV-based vegetation index than visual ratings. The significant correlations between OSAVI and the visual ratings from the study suggest that undesirable variability associated with visual assessments can be minimized with the UAV-based approach. UAV-based imagery methods had greater precision than the visual-based ratings for crop herbicide damage. These methods have the potential to replace visual ratings and aid in screening crops for herbicide tolerance.

Weed management in organic lentil (Lens culinaris Medik.) is challenging due to its poorly competitive nature with weeds and the lack of organic herbicides for field crops. In-crop mechanical weed control methods such as rotary hoeing, harrowing and inter-row cultivation, and the cultural practice of increased crop seeding rate, are established weed management tools in organic field crops. They have however not previously been combined into an integrated weed management system for weakly competitive grain crops. The objective of this research was to determine the effect of mechanical weed control (rotary hoeing, harrowing and inter-row cultivation) and crop seeding rate alone and in combination on weed suppression and yield in organic lentil. A two-factor randomized complete block design experiment was conducted at two locations in Saskatchewan, Canada in 2016 and 2017. The factors mechanical weed control methods including the rotary hoe, the harrow and inter-row cultivation and an untreated control, and seeding rates including conventional recommended and double recommended seeding rate. All treatments were tested as stand-alone or combined in a factorial fashion. Increased crop seeding rate and mechanical weed control affected response variables independently of each other. Doubling the seeding rate of lentil reduced weed biomass by 16% and increased lentil yield by 23% averaged across the four site-years. Treatments that included only a single mechanical weed control methods resulted in poor or incomplete weed suppression at all sites. Two mechanical weed control methods were required to maximize yield, and three were required to minimize weed biomass. Early weed control with a rotary hoe, followed by inter-row cultivation prior to canopy closure was the most consistent mechanical weed control treatment across the four site-years. Averaged across the four site-years, rotary hoeing + inter-row cultivation reduced weed biomass by 70%, and increased crop yield by 65%. The treatment maintained low weed populations across the longest time duration and controlled weeds in both the inter- and intra-row spaces, while minimizing crop injury. Doubling crop seeding rate and using two appropriately timed mechanical weed control methods provided weed suppression while increasing yield by 46% in a weakly competitive organic lentil crop.

Seedbank persistence in canola seeds is related to their potential to develop secondary dormancy. This can result in volunteer weed problems many years after canola production. The potential to be induced into secondary dormancy is controlled by both the canola genetics and the environment of the mother plant. However, the effect of time of harvesting on secondary dormancy potential is not known. The objective of this study was to determine the effect of harvest timing on potential to develop seed dormancy in canola. Six harvest samples were collected weekly from two canola genotypes (5440 and 5020) starting from 10 to 20% seed color change on the main stem until they were fully ripened. Freshly harvested seeds of 5440 and 5020 showed 13 and 16% primary dormancy at 32 and 33 d after flowering (DAF), respectively, but dormancy decreased with harvest timings and no dormancy was observed when seeds were fully mature (78 DAF). After dormancy induction, 10% of 5440 seeds were dormant at 32 DAF, but 94% of seeds were dormant at 78 DAF. Similarly, 70% of 5020 seeds were dormant at 33 DAF, but 90% of seeds were dormant at 68 DAF. Thus, seeds had lower potential to secondary dormancy at early development but have a high potential to secondary dormancy induction at full maturity. This study suggests that windrowing these canola genotypes at the recommended time (60% seed color change on the main stem) may reduce ability of the seed to develop secondary dormancy and thus reduce the persistence of seeds in the soil seedbank.

Cowcockle, an introduced summer annual weed of the Northern Great Plains, is being considered for domestication because of its high quality starch, cyclopeptides, and saponins. Loss of seed dormancy is one of the key desirable traits for domestication. To determine the potential for domestication of this species, an understanding of the seed dormancy and germination patterns is required. The objectives of this study were to evaluate seed dormancy in cowcockle ecotypes and determine how temperature and light affect seed dormancy. We evaluated 15 populations of cowcockle for primary dormancy by exposing them to five temperatures (5, 7.5, 10, 15, and 20 C) under two temperature regimes (constant and alternating) in both dark and light conditions. Freshly matured seeds of all the populations showed high levels of primary dormancy except ‘Mongolia’. Lower levels of dormancy at medium temperatures (10 and 15 C) and greater dormancy at low and high temperatures suggest conditional dormancy, a state at which seeds germinate over a narrower range of conditions compared to nondormant seeds. The effects of temperature regime, light, and their interaction was significant only at suboptimal (5 and 7.5 C) and supraoptimal (20 C) temperatures. Under these conditions, alternating temperatures were more effective in breaking the conditional dormancy, followed by light. The variation in optimum temperature, light, and their interactions among the cowcockle populations may be due to the plants evolving to adapt to their local environments. From a domestication perspective, the conditional dormancy in cowcockle can be observed as an evolutionary mechanism that prevents untimely germination following maturity and may not be a major obstacle for its domestication. Nomenclature: Cowcockle, Vaccaria hispanica (P. Mill.) Rauschert, VAPPY.

The challenges associated with the adoption of conservation tillage and/or low-input cropping systems, whether organic or herbicide-free, across Canada are shaped by scale, environment, and local practices. A study in eastern Canada captured the challenges of introducing low-input cropping systems in mature (20+ yr) tillage treatments in a barley/red clover/corn/soybean rotation. Each mature tillage system came with its own weed problems, but this did not affect crops such as barley and red clover, which produced similar yields across high and low input systems. However, some form of primary tillage was needed to achieve adequate weed control and yield in organic (ORG) and herbicide-free (HF) systems. The HF and ORG systems with no-till actually failed to produce a corn crop but produced soybean yields that were half or less than that for other treatments. The successful combination of conservation tillage practices and low-input systems in eastern Canada would thus appear to be crop-specific, being easier to achieve in competitive cereal crops. In western Canadian organic agriculture, tillage is practiced with low-disturbance chisel plows instead of inversion plows. However, green manure crops (summer cover crops) are often terminated with tandem discs. Both roller crimpers and mowing can successfully kill annual green manure crops such as field pea and rye, and usually result in reduced weed growth following termination. However, the lack of tillage can result in lower crop yields in wheat following green manure terminated by roller crimping compared to tillage. Challenges for no-till organic practices include perennial weed control and soil fertility. Overall, flexible crop production programs such as the former Manitoba Pesticide Free Production program and the “Agriculture raisonnéeTM” program in Québec are more likely to promote sustained environmental, economic, and social prosperity than rigid adherence to organic or no-till practices.

Legume crops are known to have low soil N uptake early in their life cycle, which can weaken their ability to compete with other species, such as weeds or other crops in intercropping systems. However, there is limited knowledge on the main traits involved in soil N uptake during early growth and for a range of species. The objective of this research was to identify the main traits explaining the variability among legume species in soil N uptake and to study the effect of the soil mineral N supply on the legume strategy for the use of available N sources during early growth. Nine legume species were grown in rhizotrons with or without N supply. Root expansion, shoot and root biomass, nodule establishment, N2 fixation and mineral soil N uptake were measured. A large interspecific variability was observed for all traits affecting soil N uptake. Root lateral expansion and early biomass in relation to seed mass were the major traits influencing soil N uptake regardless of the level of soil N availability. Fenugreek, lentil, alfalfa, and common vetch could be considered weak competitors for soil N due to their low plant biomass and low lateral root expansion. Conversely, peanut, pea, chickpea and soybean had a greater soil N uptake. Faba bean was separated from other species having a higher nodule biomass, a higher N2 fixation and a lower seed reserve depletion. Faba bean was able to simultaneously fix N2 and take up soil N. This work has identified traits of seed mass, shoot and root biomass, root lateral expansion, N2 fixation and seed reserve depletion that allowing classification of legume species regarding their soil N uptake ability during early growth.

Seed shatter of wild oat (Avena fatua L.), green foxtail [Setaria viridis (L.) P. Beauv.], wild mustard (Sinapis arvensis L.), cleavers (Galium spurium L. and G. aparine L.), wild buckwheat (Polygo...

Combine harvesters have the potential to disperse weed seeds great distances. Reducing this dispersal may be important in an integrated weed management system. The objectives of this study were to determine the distance that wild oat seeds are dispersed by a combine harvester and the effect of chaff collection on combine harvester seed dispersal. This was measured by sampling wild oat seeds at varying distances behind a combine equipped with a removable chaff collection system after it passed through a wild oat patch. Chaff collection consistently reduced the amount and distance that wild oat seeds were dispersed. This occurred because more than 74% of the total wild oat seed that were ejected from the combine were in the chaff. Because most of the chaff falls in a row directly behind the combine, chaff collection only affected dispersal in this area. In 1996, chaff collection reduced wild oat seed dispersal past the wild oat patch to less than 10 seeds m−2at 45 m, whereas without chaff collection, there was greater than 10 seeds m−2up to 145 m. At distances beyond 145 m, chaff collection had no significant effect on seed dispersal. Chaff collection may be an important tool in an integrated weed management program because it may slow weed invasions and reduce the expansion of weed patches.

In western Canada, little is known about the seedbank ecology of volunteer canola. Therefore, integrated recommendations for the management of this weed are limited. In this study, we investigated the seedbank persistence and seedling recruitment of two spring canola genotype groups with different secondary seed dormancy potentials under contrasting tillage systems. The study was conducted at two locations with different soils in the Mixed Moist Grassland ecoregion of Saskatchewan. A single cohort seedbank was established in 1999 and was followed for 3 yr in successive wheat crops. In a separate laboratory study, the six canola genotypes examined were classified as those with high and those with medium potentials for the development of secondary seed dormancy (HD and MD, respectively). After one, two, and three winters, maximum persistence of 44, 1.4, and 0.2% of the original seedbank was observed among the treatments, respectively. In 2001, HD canola genotypes tended to exhibit 6- to 12-fold greater persistence than MD canola genotypes, indicating lower seedbank mortality in HD canola. Seedling recruitment of HD canola also was higher than MD canola when differences were observed between these genotype groups. Therefore, long-term seedbank persistence of canola can be reduced by growing genotypes with low inherent potential for the development of secondary seed dormancy. The proportion of persisting seeds tended to be higher under conventional tillage than under zero tillage because of lower seedbank mortality, but no clear distinction in seedbank persistence in terms of absolute time could be made between these two tillage systems. Volunteer canola seedling recruitment followed the pattern of a typical summer-annual weed, where seedling emergence was observed only during May and June.